PLEASE SCROLL DOWN FOR ARTICLE

This article was downloaded by: [Casale, Achille]On: 20 July 2010Access details: Access Details: [subscription number 922693462]Publisher Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Italian Journal of ZoologyPublication details, including instructions for authors and subscription information:http://www.informaworld.com/smpp/title~content=t741771159

The third instar larva of Speomolops sardous Patrizi, 1955, a cave-dwellingmolopine beetle endemic to Eastern Sardinia, with notes on its habitat(Coleoptera, Carabidae)A. Casalea; A. Di Giuliob; P. Marciaa; A. Molinuc

a Dipartimento di Zoologia e Genetica Evoluzionistica, Università di Sassari, Sassari, Italy b

Dipartimento di Biologia, Università degli Studi 'Roma Tre', Roma, Italy c Istituto per lo Studio degliEcosistemi, Consiglio Nazionale delle Ricerche, Sede di Sassari, Sassari, Italy

First published on: 02 June 2010

To cite this Article Casale, A. , Di Giulio, A. , Marcia, P. and Molinu, A.(2010) 'The third instar larva of Speomolopssardous Patrizi, 1955, a cave-dwelling molopine beetle endemic to Eastern Sardinia, with notes on its habitat (Coleoptera,Carabidae)', Italian Journal of Zoology, 77: 2, 159 — 167, First published on: 02 June 2010 (iFirst)To link to this Article: DOI: 10.1080/11250000903015182URL: http://dx.doi.org/10.1080/11250000903015182

Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf

This article may be used for research, teaching and private study purposes. Any substantial orsystematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply ordistribution in any form to anyone is expressly forbidden.

The publisher does not give any warranty express or implied or make any representation that the contentswill be complete or accurate or up to date. The accuracy of any instructions, formulae and drug dosesshould be independently verified with primary sources. The publisher shall not be liable for any loss,actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directlyor indirectly in connection with or arising out of the use of this material.

ISSN 1125-0003 print/ISSN 1748-5851 online © 2010 Unione Zoologica ItalianaDOI: 10.1080/11250000903015182

Italian Journal of Zoology, June 2010; 77(2): 159–167

TIZOThe third instar larva of Speomolops sardous Patrizi, 1955, a cave-dwelling molopine beetle endemic to Eastern Sardinia, with notes on its habitat (Coleoptera, Carabidae)

Larva of the cave-dwelling carabid beetle Speomolops sardousA. CASALE1*, A. DI GIULIO2, P. MARCIA1, & A. MOLINU3

1Dipartimento di Zoologia e Genetica Evoluzionistica, Università di Sassari, Sassari, Italy, 2Dipartimento di Biologia, Università degli Studi ‘Roma Tre’, Roma, Italy, and 3Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Sede di Sassari, Sassari, Italy

(Received 22 October 2008; accepted 22 April 2009)

AbstractWhile the larval morphology and peculiar behaviour of several epigean species of molopine beetles (Carabidae: Pterostichini:Molopina) have been known for several years, we know very little about the specialized subterranean species of some generaendemic to the Tyrrhenian area, including Molopidius Jeannel, 1942, Oscadytes Lagar, 1975, Henrotius Jeannel, 1953,Zariqueya Jeannel, 1924, and Speomolops Patrizi, 1955. In this paper, we describe and illustrate the third instar larva ofSpeomolops sardous Patrizi, 1955, a specialized troglobitic beetle endemic to eastern Sardinia. Diagnostic features includethe lack of stemmata; the presence of a longitudinal, membranous furrow along the dorsal surface of the mandibles; a smallretinaculum; and a uniquely shaped seta FR8. We compare the morphology of this larva with some of its close relatives inthe genera Molops Bonelli, 1810, Tanythrix Schaum, 1858, and Typhlochoromus Moczarsky, 1913. Some data are also givenabout the habitat and bionomics of the larval stages of Speomolops sardous, which show an obligate, fossorial behaviour intosandy soils in deep zones of large, periodically flooded subterranean systems.

Keywords: Coleoptera, Carabidae, Speomolops sardous, larval morphology, habitat

Introduction

The monophyly of the pterostichine subtribe Mol-opina Bonelli, 1810 is only moderately supported(Brandmayr & Zetto Brandmayr 1994; Düring &Brückner 2000). Diagnostic characters of the adultsinclude the absence of discal setiferous punctures onelytra, the presence of a basal keel on the seventh (orsixth) elytral interval (in some taxa, however,reduced or absent), the lack of secondary sexualcharacteristics in the seventh abdominal segment inthe male, and the presence of a membranous bandat the base of the first antennomere in the larvae.The features of female genitalia do not provideadditional informative data on the monophyly ofMolopina (Giachino & Sciaky 1991).

From the molecular point of view, however, theclade “Molopina”, including the Euro-Mediterraneangenera Molops Bonelli, 1810, Tanythrix Schaum,

1858, Typhlochoromus Moczarski, 1913 (treated asdistinct genera: Vigna Taglianti 2005), PercusBonelli, 1810, Abax Bonelli, 1810, and some subter-ranean taxa cited below, but excluding StyracoderusChaudoir, 1874 (an Iberian genus currently attrib-uted to this subtribe: Serrano 2003) and all generafrom the southern hemisphere (South Africa, Mada-gascar and Australia, attributed to Molopina insome contributions: see, among others, Jeannel1948), seems to be strongly supported (Casale &Ribera, in press).

Interesting biogeographic patterns shown by spe-cies in some genera of Molopina have been stressed inseveral papers (Baroni Urbani et al. 1978; Brückner& Mossakowski 2006), and are particularly evident insome subterranean genera. Among these, the mono-typic genus Speomolops Patrizi, 1955 (with the speciessardous Patrizi, 1955) is known only from a few cavesin Triassic limestone massifs of Central–Eastern

*Correspondence: A. Casale, Dipartimento di Zoologia e Genetica Evoluzionistica, Via Muroni 25, 07100 Sassari, Italy. Email: casale@uniss.it

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

160 A. Casale et al.

Sardinia. From the morphological point of view,Speomolops is closely related to the genera MolopidiusJeannel, 1942, Oscadytes Lagar, 1975, HenrotiusJeannel, 1953, Zariqueya Jeannel, 1924, the speciesof which are endemic to the Tyrrhenian area(Southern Iberian peninsula, Pyrenees, Baleares).This distribution pattern is currently interpreted asthe result of an ancient vicariant event produced bythe tectonic, Miocene drift of the West Mediterra-nean micro-plates (Vigna Taglianti 1982; Casale &Vigna Taglianti 1996).

Representatives of some genera within Molopinaare highly notable for parental care, described morethan one century ago (Bargagli 1874), and which israther unusual among carabids (Brandmayr & ZettoBrandmayr 1979; Brandmayr 1992; Kavanaugh1998).

Nevertheless, and in spite of a rich literature onthese beetles, including descriptions of several larvalstages, nothing was known so far about the pre-imaginal stages of the subterranean taxa of the WesternMediterranean area, their development, habitatchoice, and behaviour.

In this contribution, the authors describe the mor-phological features of the third instar (mature) larvaof Speomolops sardous Patrizi, 1955, compared withthe larval morphology of related species of Molopina(genera Molops, Tanythrix and Typhlochoromus), withnotes on the behaviour and habitat of this highlyspecialized troglobitic species.

Materials and methods

Material examined: five larvae (one second and fourthird instars) of Speomolops sardous, preserved in70% ethyl alcohol: “Italy, Central–Eastern Sardinia,Bue Marino cave near Cala Gonone, Dorgali(Nuoro), Sardinian Speleological cadastre: 12 SA/NU, 40° 14′ 51″ N, 9° 37′ 29″ E, 10/9/2006”.Samples were collected during one of the severalbio-speleological expeditions organized in last yearsby some of the authors (A.C., A.M., P.M.), and byother bio-speleologists working in this area (seeCasale et al. 2008). The identification of larvae wasstated ex societate imaginis. This material is presentlydeposited in the collections of the Department ofZoology, University of Sassari (Italy).

Two third instar larvae were cleared in 10%KOH, dehydrated in ethanol, and mounted onmicroscope slides in Canada Balsam. Another thirdinstar larva was dehydrated in ethanol, cleared inxylene, and mounted as a dried specimen on apinned card. The second instar larva was criticalpoint-dried, sputtered with gold, and analysed with

a Philips XL30 scanning electron microscope(L.I.M.E. laboratory, University “Roma Tre”,Rome). This larva was used for comparison, but isnot described in this work.

The specimens have been examined and drawnwith stereomicroscopes Wild M5 and OlympusSZX16, and for details with Zeiss Orthoplan andBX51, all equipped with drawing tubes.

Measurements were made by analysing picturestaken by a digital camera Color View II (Soft Imag-ing System) by Cell* (software for life sciencesmicroscopy, Olympus Cell* Family, Soft ImagingSystem). The main measures considered, with acro-nyms, are the following: total body length (TL),from anterior margin of frontale (at the level ofnasale) to apex of abdominal segment X (pygidium);head maximum length (HL), from anterior marginof epistoma (apex of nasal teeth) to base of stem ofepicranial suture; head maximum width (HW);length of pronotum (LP); width of pronotum (WP);length of mesonotum (LM); width of mesonotum(WM); length of urogomphi (UL).

For general terminology of larval structures werefer to Lawrence (1991). Notation of primary setaeand pores follows the system of Bousquet andGoulet (1984) and Bousquet (1985).

Results

Description of the third instar larva of Speomolops sardous

Habitat. Larvae were collected in deep zones of thecave, about 2 km from the entrance into sandy, humidsoil on the banks of one of the internal subterraneanlakes, where many adult individuals were walking.

Diagnosis. This larva can be distinguished fromother molopine larvae for the peculiar combinationof the following morphological characters: stemmataabsent; mandibles with a dorsal, longitudinal, mem-branous furrow; retinaculum very small; seta FR8sickle-shaped, widely flattened and expanded.

Measurements. TL: 8.58 mm; HL: 0.77 mm; HW:0.92 mm; LP: 1.06 mm; WP: 1.18 mm; LM: 0.69mm; WM: 1.18 mm; UL: 1.25 mm.

Habitus and coloration. Body soft, slender, cylindri-cal, subparallel sided, weakly sclerotized (Figures 1and 2). Colour pale, with cephalic capsule and man-dibles more sclerotized, reddish-brown; head append-ages, legs and urogomphi light brown; sclerites ofthorax and abdomen pale, whitish to light brown.

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

Larva of the cave-dwelling carabid beetle Speomolops sardous 161

Microsculpture. Sclerotized structures smooth, mem-branous areas of body rugose. Anterior part of prono-tum (Figure 12) distinctly corrugated longitudinally.

Setae of head and body longitudinally corrugatedexcept for setae of urogomphi with irregular transversalpattern.

Figures 1–3. Speomolops sardous, larval and adult features: 1, third instar larva, habitus, dorsal aspect; 2, idem, alive individual on sandysoil near an inner lake of Bue Marino cave; 3, adult individual walking on sandy soil.

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

162 A. Casale et al.

Chaetotaxy. Frontoclypeolabrum (Figures 4, 7) with1 pair of secondary setae on adnasalia internal andparallel to FR9; FR8 sickle-shaped, widely flattenedand expanded; FR3 extremely small; FR4 slightlyposterior to the level of pore FRc; FR10 and FR11

peg-like. Parietale (Figures 4–6) without secondarysetae; PA4, PA8 and PA14 very small; PA5 posteriorto cervical groove. Antennae (Figure 8) with anten-nomere I with 1 secondary seta; II with 2 secondarysetae. Mandible (Figure 9) without secondary setae.

Figures 4–11. Speomolops sardous, third instar larva: 4, head, dorsal view; 5, head, ventral view; 6, head, lateral view; 7, anterior margin offrontoclypeolabrale; 9, right mandible, dorsal view; second instar larva: 8, left antenna, dorsal view; 10, left maxilla, dorsal view; 11,labium, ventral view.

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

Larva of the cave-dwelling carabid beetle Speomolops sardous 163

Maxilla (Figure 10) with setal group gMX on stipescomposed of approximately 50 small setae; onestrong, spiniform, short seta on outer margin, morebasal than MX2; seta MX3 about twice as long asMX2; MX10 very small; palpomere IV with smallmedial secondary seta, six longitudinal subapical digi-tiform sensilla and apical sensorial area composed ofseveral small papillae. Prementum (Figure 11) with-out secondary setae; labial palpomere II with onevery small secondary seta, medially directed, and api-cal sensorial area composed of several small papillae.Pro-, meso- and metanotum (Figure 12) withoutsecondary setae. Coxa, trochanter and femur (Figure13) with setae spine-like; no secondary setae present.Abdominal tergites (Figure 1) without secondarysetae. Median sternites (Figure 14) with two pairs ofsecondary setae. Hypopleurites (Figure 14) withthree secondary setae. Epipleurites (Figure 14) withtwo secondary setae. Urogomphi (Figure 15) withtwo ventral secondary setae each, one subapical andone medial; one basal secondary seta often presentlaterally. Pygidium without secondary setae.

Head. Cephalic capsule (Figures 4–6) slightly trans-verse (HW/HL = 1.18), prognathous, subparallelsided, only slightly curved; maximum width at levelof setae PA6; occipital foramen dorsally displaced.Frontoclypeolabrum (Figures 4, 7) shield-like,slightly elongate (L/W = 1.2), with surface mediallyconvex and laterally concave, anteriorly sloped;frontoclypeolabral length about 6.5 times longerthan basal stem of epicranial suture, anterior frontalarms straight or only slightly sinuate; frontoclypeola-brum anteriorly produced into two subtriangularteeth (Figure 7), apically rounded, widely separatedby a deep, V-shaped, incisure; frontoclypeolabraleconvex at the level of the teeth; adnasalia pointedand slightly protruding; epipharynx with preoralfilter apparatus, composed by two lateral tufts ofverticillate hairs projecting towards the mouth and amedial tuft of plumy hair structures, partiallybranched, anteriorly directed. Parietale (Figures 4–6)without any trace of stemmata; ocular grooveabsent; cervical groove wide, slightly extendedventrally; walls of parietale medially fused, forming a

Figures 12–15. Speomolops sardous, third instar larva: 12, pro- and mesonotum, dorsal view; 14, abdominal segment III, ventral view; 15,abdominal apex with urogomphi, dorsal view; second instar larva: 13, foreleg, posterior view.

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

164 A. Casale et al.

complete gular suture (Figure 5). Antennae (Figures4, 6, 8) four-jointed, about as long as mandibles,forwardly directed; antennomere I longer thanothers, with sub-basal membranous belt; II half longthan I; III slightly shorter than I; IV half long thanIII; III with conical sensorial appendage, about halfas long as IV. Mandibles (Figure 9) slender, sickle-shaped, distinctly upcurved, about 2.5 times as longas their basal width, strongly curved along lateralmargin, with a dorsal, longitudinal, deep membra-nous furrow, extending along basal third; penicilluspresent; cutting edge smooth, regularly concave; ret-inaculum very small, subtriangular, basally directed,with sharp apex. Maxilla (Figure 10) with elongateand slender stipes curved inwards, about three timesas long as their basal width; maxillary palpomere IIIlonger than others gradually decreasing in lengthfrom III to I; IV slender, cylindrical, about as long asI; lacinia (Figure 10) small and pointed, inwardlydirected; galea slender divided into two joints,galeomere II slender, slightly longer than I. Labium(Figure 11) with subquadrate prementum; ligula notapparent; labial palpomere I slightly shorter andwider than II.

Thorax. Pro-, meso- and metanotum (Figures 1,12) of the same width, with longitudinal ecdysialline well marked. Pronotum about 1.5 times longerthan meso- and metanotum, subtrapezoidal, aboutas long as wide, with maximum width at base.Meso- and metanotum widely transverse, about twotimes wider than long.

Spiracles. Thoracic and abdominal spiracles sub-equal, very small (35–40 μm), annular, not protrud-ing; mesothoracic spiracles positioned anterolaterallyon mesopleura. Abdominal spiracles slightly dorsal toepipleurites.

Legs. Five-jointed (Figure 13), foreleg distinctlyshorter than meso- and hindleg. Coxa subcylindri-cal, longer than other joints; trochanter and femursubequal, 2/3 than coxa; tibia and tarsus subequal,2/3 than femur; trochanter obliquely truncate andventrally expanded at apex; claws strong, subequal,only slightly curved and pointed at apex.

Abdomen. Cylindrical, subparallel sided, with seg-ments I–VII subequal, subrectangular (Figure 1);VIII slightly narrower; IX suboval, bearing a pair ofelongate urogomphi (Figures 1, 15). Tergites trans-verse, with medial longitudinal suture (Figure 1),and sides slightly rounded. Epi- and hypopleuriteselongate, subrectangular (Figure 14). Sternal areaof segments I–VII composed by two small anterior

sternites, one suboval median sternite, two poste-rior inner sternites and two lateral outer sternites(Figure 14); medial sternite and inner and outersternites fused into a unique sternal sclerite on ster-num VIII, incorporating also anterior sclerites onsternum IX. Urogomphi fix, elongate, slender,apically upcurved, not segmented, with distinctsetiferous nodes (Figure 15). Pygidium elongate,cylindrical, about as long as segment IX, with dorsaland ventral sclerites not fused.

Discussion

The discovery of larval stages of highly specializedtroglobitic carabids is a rare event, even in caveswhere adult individuals are very abundant (Casale &Marcia 2008). This fact induced some bio-speleologiststo suspect that most specialized troglobitic carabidsmay have a contracted lifecycle, reduced to a singlelarval stage which does not eat, as in some cholevidbeetles of the tribe Leptodirini. On the contrary, ithas been demonstrated that many carabid beetlesthat are highly adapted to a subterranean environ-ment have running, predaceous larvae, morphologi-cally very similar to those of epigean species (Casaleet al. 1998). This is the case of the larva of Speomol-ops sardous here described. In spite of the fact thatboth adults and larvae display marked modificationsfor life in caves (blindness, general weak sclerotiza-tion, pale colour body), the larval features are in fullagreement with the general morphological form ofepigean pterostichine larvae, other than those ofZabrini and Abacetini (Emden 1942; Jeannel 1942;Sharova 1958; Thompson 1979a,b; Bousquet 1985;Makarow 1994): frontoclypeolabrum prominent;cervical groove present (occipital groove of originalauthors), wide; mandibles elongate and slender,with retinaculum inserted at middle length of theinner side; legs each with two, subequal tarsal claws.

Among the known Molopina larvae, species of thegenera Percus Bonelli, 1810 and Abax Bonelli, 1810,which are closely related to each other, are easily rec-ognizable for having subovate head, frontoclypeola-brum with anterior margin straight or gently curved,without deep incisure in the middle, and segmentedurogomphi (Beer & Binaghi 1937; Lumaret 1971;Arndt 1989). Larval features of Speomolops sardousare similar to those of some species of genera Mol-ops, Tanythrix and Typhlochoromus, described byCerruti (1941) and Leonardi (1969). In these lar-vae, the frontoclypeolabrum shows two evidentouter lobes (adnasalia) and two inner lobes widelyseparated by a deep furrow; in Typhlochoromus thefurrow is markedly deep as in Speomolops, but it isnarrower and U-shaped. The sub-basal membranous

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

Larva of the cave-dwelling carabid beetle Speomolops sardous 165

belt in antennomere I is present in all molopine gen-era cited above. Whereas the adnasalia in Molops,Tanythrix and Typhlochoromus have either six orseven setae, in Speomolops there are only three setae,and seta FR8 is unique in being sickle-shaped,widely flattened and expanded.

Unfortunately, we did not obtain first instar larvaeof Speomolops sardous, which would have beenimportant for a comparison of the primary chaeto-taxy with other Molopina genera. However, larvalstages of Speomolops sardous are markedly distinctfrom all other molopine larvae for the peculiarcombination of the following features: lacking stem-mata, and for having mandibles with a dorsal, mem-branous furrow, a retinaculum very reduced in size,and a maxilla with a setal group gMX on stipescomposed of many small setae.

Conclusion

The phylogenetic position of the genus Speomolopsamong Molopina, and its interesting chapter in thehistory of the Mediterranean insect fauna, have beendebated several times. In particular, based on adultmorphology confirmed by molecular data (Ribera,personal communication), it is possibly the adelpho-taxon of the Iberian genus Zariqueya Jeannel, 1924,or the Balearic genus Henrotius Jeannel, 1953 (VignaTaglianti 1982; Casale & Vigna Taglianti 1996).

Larval features described here do not support (orreject) this hypothesis, since no other troglobiticMolopina species are unknown so far. Nevertheless,the discovery of larval stages of S. sardous is veryinformative from several different bio-speleologicalpoints of view. In particular, it confirms that inhighly specialized, troglobitic carabid beetles thelarval stage is not reduced to a single instar, butrather is very similar to closely related epigean taxa.In fact these larvae are in agreement with the “soilpore explorer” type proposed by Zetto Brandmayret al. (1998) and with the “stratobionts-runners, ofthe cryptobionts-series” of Sharova (2008). Further-more, several characteristics of Speomolops larvae,such as the lack of stemmata, weak sclerotization,unspecialized mandibles, moderately long, unarticu-lated urogomphi, are well-known adaptations to lifein caves. On the other hand, some of these modifica-tions, such as the absence of stemmata, are alsoknown in subterranean larvae of species withepigean adults belonging to different groups ofcarabids, such as the “ant killer” Siagona europaea,the adults of which have functional eyes (ZettoBrandmayr et al. 2007).

In Bue Marino cave, adults of Speomolops sardousare numerous. They are often found walking on

sandy, humid soil on the banks of the subterraneanlakes in the inner parts of the cave. On the contrary,the larva of this species was searched in vain, formore than 30 years, by one of the authors (A.C.)(Casale 1972; Casale et al. 2008). Finally, duringonly one visit on 10 September 2006, five larvaewere collected 20–30 cm below the surface of asandy bank near one of the freshwater lakes whereadults were very abundant. The larvae were neverobserved walking on the surface. This demonstratesthat the larva of this species has obligate under-ground, fossorial behaviour, in sandy soil periodi-cally flooded by the subterranean river of the cave.That this larva was finally discovered confirms theimportance of the periodical monitoring pro-gramme, carried out for three years by one of theauthors (P.M.) and presently in progress, for theconservation in Bue Marino cave of this, and othersubterranean species, endemic to this area ofSardinia. This cave is, in fact, one of the mostattractive show caves of the island, visited by nearly80,000 people each year. Adult Speomolops sardouswere found only in private areas of the cave, notopen to tourists, and they seem to have a rather highdispersal power in deep crevices (the species isknown by a few specimens or remains from someother Sardinian caves: Casale et al. 2008). However,the larval stages show an obligate, fossorial way oflife in sandy soils within large, undisturbed areas ofthe cave, which are periodically flooded. Any furtherhuman impact on these environments, such asmodifications induced by ecotourism, may have adestructive effect on the survival of this endemic,relict species.

Acknowledgements

For the invaluable speleological support in the field,the authors are particularly indebted to GiuseppeGrafitti (Sassari), Enrico Lana (Torino), MauroMucedda (Sassari), Carlo Onnis (Cagliari), VirginiaDenanni (Sassari) and many other speleologists whocontributed to the biological exploration of the BueMarino cave, and for the facilities in visits andexploration of this cave to Fabio Sagheddu and theAtlantika team (Dorgali).

For communication of data and bibliographicreferences, we are very grateful to Erik Arndt(Anhalt University of Applied Sciences, Bernburg,Germany), and to Pietro Brandmayr and TulliaZetto (Università della Calabria, Arcavacata diRende, Italy).

We are also indebted to Wendy Moore (Universityof Arizona, Tucson, USA) for the linguistic revisionof the English text, and to three anonymous reviewers,

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

166 A. Casale et al.

who highly improved the original version of themanuscript.

Funds for this research were met with a grant from aPRIN project of the Italian Ministry of University andResearch, from the University of Sassari, and theINTERREG3 project Sardinia–Corsica–Tuscany.

References

Arndt E. 1989. Die Larven der mitteleuropaischen Abax s. str.-Arten (Coleoptera, Carabidae: Pterostichini). Beitraege zurEntomologie 39:255–270.

Bargagli P. 1874. Cenni biologici su due specie di Percus. Bullet-tino della Società entomologica italiana 6:27–31.

Baroni Urbani C, Ruffo S, Vigna Taglianti A. 1978. Materiali peruna biogeografia italiana fondata su alcuni generi di ColeotteriCicindelidi, Carabidi e Crisomelidi. Memorie della Societàentomologica italiana 56 (1977):35–92.

Beer S, Binaghi G. 1937. Percus dejeani Dj. (Col. Car.).Descrizione della larva. Bollettino di Zoologia agraria eBachicoltura, Torino 7 (1936):3–7.

Bousquet Y. 1985. Morphologie comparée des larves de Pterosti-chini (Coleoptera: Carabidae): Descriptions et tables détermi-nation des espèces du nord-est de l’Amérique du nord. Revued’écologie et de systématique 112:191–251.

Bousquet Y, Goulet H. 1984. Notation of primary setae andpores on larvae of Carabidae (Coleoptera: Adephaga).Canadian Journal of Zoology 62:573–588.

Brandmayr P. 1992. Short review of presocial evolution in Coleop-tera. Ethology, Ecology and Evolution, Special Issue 2:7–16.

Brandmayr P, Zetto Brandmayr T. 1979. The evolution of paren-tal care phenomena in Pterostichini, with particular referenceto the genera Abax and Molops. In: den Boer PJ, Thiele HU,Weber F, editors. On the evolution of behaviour in Carabidbeetles. Miscellaneous Papers, 18, LandbouwhogeschoolWageningen, pp. 35–49.

Brandmayr P, Zetto Brandmayr T. 1994. The evolutionary historyof the genus Abax (Coleoptera, Carabidae). In: Desender K,Dufrêne M, Loreau M, Luff ML, Maelfait J-P, editors. Carabidbeetles: Ecology and evolution. Dordrecht: Kluwer, pp. 19–24.

Brückner M, Mossakowski D. 2006. Phylogeny of the genus Per-cus (Coleoptera: Carabidae) – nuclear genes and the basalsplits. In: Serrano J, Koivula M, Lövei G, editors. Proceedingsof the XII European Carabidologists’ Meeting. EntomologicaFennica 17:195–199.

Casale A. 1972. Visione d’insieme del complesso ecologico e fau-nistico della Grotta del Bue Marino. Bollettino della SocietàSarda di Scienze naturali 10:3–28.

Casale A, Grafitti G, Lana E, Marcia P, Molinu A, Mucedda M,Onnis C, Stoch F. 2008. La Grotta del Bue Marino: Cin-quanta anni di ricerche biospeleologiche in Sardegna. Atti XXCongresso Nazionale di Speleologia (Iglesias, 2007), Memo-rie dell’Istituto italiano di Speleologia, Bologna (S. II)XXI:197–209.

Casale A, Marcia M. 2008. Larval morphology of Sardaphaenopsadelphus Casale, 2004, a highly specialized troglobitic beetleendemic to Sardinia (Coleoptera, Carabidae). SubterraneanBiology 5:35–42.

Casale A, Ribera I. (in press). Are Molopina of the Euro-Mediterranean region related to the Madagascan, South Africanand Australian Pterostichini? (Coleoptera, Carabidae). Biogeo-graphia (n.s.).

Casale A, Vigna Taglianti A. 1996. Coleotteri Carabidi di Sardegnae delle piccole isole circumsarde, e loro significato biogeografico

(Coleoptera, Carabidae). Biogeographia, Lavori della Societàitaliana di Biogeografia (n.s.) 18 (1995):391–427.

Casale A, Vigna Taglianti A, Juberthie C. 1998. ColeopteraCarabidae. In: Juberthie C, Decu V, editors. Encyclopaediabiospeologica, 2. Moulis-Bucarest, pp. 1047–1081.

Cerruti M. 1941. Larva di Molops ovipennis ssp. istrianus G. Müll.(Col.). Mitteilungen der Münchner Entomologischen Gesells-chaft e. V., XXXI (XXXII):938–943+1 plate.

Düring A, Brückner M. 2000. The evolutionary history of thetribe Molopini: A first molecular approach. In: Brandmayr P,Lövei G, Zetto Brandmayr T, Casale A, Vigna Taglianti A,editors. Natural history and applied ecology of Carabid beetles.Sofia-Moscow: Pensoft. pp. 1–4.

Emden FI van 1942. A key to the genera of larval Carabidae(Col.). Transactions of the Royal Entomological Society,London 92:1–99.

Giachino PM, Sciaky R. 1991. Valore sistematico delle strutturegenitali femminili in Pterostichinae (Coleoptera: Carabidae).Atti XVI Congresso nazionale italiano di Entomologia, 23/18settembre 1991, Bari-Martina Franca (TA), pp. 885–892.

Jeannel R. 1948. Coléoptères Carabiques de la Région Malgache(Deuxième Partie) Faune de l’Empire Français, X, Editionsdu Muséum. pp. 373–765.

Kavanaugh DH. 1998. Field observations confirming brood carein Percus passerinii in the Tuscan Apennines, Italy (ColeopteraCarabidae). Bollettino della Società entomologica italiana130:55–63.

Lawrence JF. 1991. Order Coleoptera (general discussion, familykey, various family treatments). Immature insects, 2. Dubu-que, Iowa: Kendall/Hunt. pp. 144–658.

Leonardi C. 1969. Descrizione delle larve di Molops senilis Schaum eTyphlochoromus stolzi (Mocz.) e note sulla sistematica larvale diquesti due generi di Pterostichini (Col. Carabidae). Annali delMuseo civico di Storia naturale, Genova 77:609–625.

Lumaret J-P. 1971. Stades larvaires de Percus (Pseudopercus) nav-aricus Dejean (Col. Pterostichidae). Bulletin de la Sociétéentomologique de France 76:55–60.

Makarow KV. 1994. A key to the genera of the Ground-beetle larvae(Coleoptera, Carabidae) of the Palearctic region. Bollettino delMuseo regionale di Scienze naturali, Torino 12:221–254.

Patrizi S. 1955. Nuovo genere e nuova specie di Pterostichide troglo-bio della Sardegna orientale (Coleoptera Caraboidea, Fam. Pter-ostichidae, Trib. Molopini). Fragmenta Entomologica 2:29–40.

Serrano J. 2003. Catalogo de los Carabidae (Coleoptera) de laPeninsula Iberica. Zaragoza: Monografias S.E.A., 9, 130 pp.

Sharova IK. 1958. The larvae of Carabidae, beneficial and nox-ious to agriculture. Scientific notes of Moscow State Pedagog-ical Institute 124:1–165 [in Russian].

Sharova IK. 2008. Adaptive radiation of carabid larvae (Coleoptera,Carabidae). In: Penev L, Erwin T, Assmann T, editors. Back tothe roots and back to the future. Towards a new synthesisamongst taxonomic, ecological and biogeographical approachesin carabidology. Proceedings of the XIII European Carabidolo-gists Meeting, 20–24 August 2007, Blagoevgrad. Sofia-Moscow:Pensoft. pp. 285–304.

Thompson RG. 1979a. A systematic study of larvae in the tribesPterostichini, Morionini, and Amarini (Coleoptera: Carabi-dae). Bulletin of the Agricultural Experimental Station,Arkansas University, Fayetteville 837:1–105.

Thompson RG. 1979b. Larvae of North American Carabidaewith a key to the tribes. In: Erwin TL, Ball GE, WhiteheadDR, editors. Carabid beetles, their evolution, natural history, andclassification. The Hague: W. Junk Publishers. pp. 209–291.

Vigna Taglianti A. 1982. Le attuali conoscenze sui ColeotteriCarabidi cavernicoli italiani. Lavori della Società Italiana diBiogeografia (n.s.) 7 (1978):339–430.

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010

Larva of the cave-dwelling carabid beetle Speomolops sardous 167

Vigna Taglianti A. 2005. Checklist e corotipi delle specie di Carabi-dae della fauna italiana. Appendice B. In: Brandmayr P, Zetto T,Pizzolotto R, editors. I Coleotteri Carabidi per la valutazioneambientale e la conservazione della biodiversità. Manuale opera-tivo. APAT, Manuali e linee guida, 34. pp. 186–225.

Zetto Brandmayr T, Giglio A, Marano I, Brandmayr P. 1998. Mor-phofunctional and ecological features in carabid (Coleoptera) lar-vae. In: Ball GE, Casale A, Vigna Taglianti A, editors. Phylogeny

and classification of Caraboidea. Atti XX International Congressof Entomolgy (Florence, 1996), Museo Regionale di ScienzeNaturali, Torino. pp. 449–490.

Zetto Brandmayr T, Mazzei A, Talarico F, Giglio A, Bauer T,Brandmayr P. 2007. The larva of Siagona europaea Dejean,1826: Morphology and collecting technique for a subterra-nean blind “running ant killer” (Coleoptera, Carabidae).Italian Journal of Zoology 74:239–245.

Downloaded By: [Casale, Achille] At: 18:22 20 July 2010